Many of the bolded characters in the characterization above are apomorphies of subsets of streptophytes along the lineage leading to the embryophytes, not apomorphies of crown-group embryophytes per se.

All groups below are crown groups, nearly all are extant. Characters mentioned are those of the immediate common ancestor of the group, [] contains explanatory material, () features common in clade, exact status unclear.

Age. The age of this node is estimated at (105-)84(-58) m.y. (Lemaire et al. 2011b); K. Bremer et al. (2004) suggested an age of ca 113 m.y., Magallón and Castillo (2009: c.f. position of Paracryphiales) an age of ca 92 m.y., Magallón et al. (2015) an age of around 85.6 m.y., Naumann et al. (2013) an age of around 75.2 m.y., and Beaulieu et al. (2013a: 95% HPD) an age of (101-)91(-79) m.y.; around 92 or 86.4 m.y. are ages in Nylinder et al. (2012: suppl.) and ca 124 m.y.a. in Nicolas and Plunkett (2014).

Divergence & Distribution. Initial diversification of this clade probably occurred in the southern hemisphere, but diverse clades in it like Apiaceae "coincide" with more recent movements to the north (Beaulieu et al. 2013a).

For the fossil record of the order, see Martínez-Millán (2010) and Nicolas and Plunkett (2014).

Note: Possible apomorphies are in bold. However, the actual level at which many of these features, particularly the more cryptic ones, should be assigned
is unclear. This is partly because many characters show considerable homoplasy, in addition, basic information for all too many is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there is the not-so-trivial issue of how ancestral states are reconstructed (see above).

Evolution.Divergence & Distribution. Apiales contain ca 2.4% eudicot diversity (Magallón et al. 1999). Although Magallón and Sanderson (2001) estimated that diversification in Apiales had increased, as Nicolas and Plunkett (2014) correctly point out, the bulk of the diversity in Apiales is in Apiaceae.

Nicolas and Plunkett (2014: conclusions not affected by the position of Pennantiaceae) suggest that Pennantiaceae, Torricelliaceae, Griseliniaceae, and a clade representing the rest of the order are ancient (Cretaceous) and possibly of East Gondwanan origin, although East Asia is also involved. They thought that vicariance might be involved in some of the distributions, e.g. of the three genera of Torricelliaceae, etc..

Thinking about morphological evolution is particularly difficult here because of incomplete knowledge of and extensive variation in the characters of interest. The basal pectinations are genera that have transseptal bundles in common, as well as some rays over 10 cells
wide and with square or upright cells (Noshiro & Baas 1998; Baas et al.
2000). Pennantia, with a superior ovary, may also be placed here (see below). Unfortunately, corolla
initiation, pollen cell number, and many other characters are unknown for
these taxa, although some information for the ex-Cornalean genera can be found in Patel (1973),
Philipson (1967), and Philipson and Stone (1980). Pennantia has similar (plesiomorphic) wood anatomy, rather like Griseliniaceae but unlike that of other other Apiales (Lens et al 2008a); this suite of characters is scored as having two origins in Apiales... Griseliniaceae and
Torricelliaceae both have the iridoid griselinoside, transseptal bundles in the ovary, and the abaxial carpel alone is fertile, the single
ovule being apotropous. Pennantia has a superior ovary (and sometimes a thick, disciform, sessile stigma). In particular, how the gynoecium of Pennantia is interpreted (summarized in Kårehed 2003a) has many implications for character evolution. Here the gynoecium is interpreted as being tricarpellate, although it appears to be single ("pseudomonomerous"), and the carpel that is fertile is abaxial (see also Chandler & Plunkett 2004; Plunkett et al. 2004c for carpel number).

Erbar and Leins (1988a,
1996a, 2004) suggested that the nectary "disc" of Apiaceae and Araliaceae is a carpellary
flank nectary displaced by intercalary growth, and that the axile placentation and inferior ovary of
these two families is easily related to the parietal placentation and superior ovary of
Pittosporaceae. The ovary of Pittosporaceae has a short basal zone with separate
loculi, the ovules being borne above this zone - essentially the same position
in which the ovules of Apiaceae and Araliaceae are found. However, gynoecial development, etc., in the whole Apiales badly needs reinvestigation, since apart from Pennantiaceae (perhaps sister to the rest of the order), nearly all Apiales have inferior ovaries. Furthermore, clades with with superior and inferior ovaries interdigitate in most other campanulid orders.

Further complicating the issue, sister to the [Apiales [Paracryphiales + Dipsacales]] clade is Bruniales, a small but morphologically very heterogeneous clade, while within Apiales, Pittosporaceae are florally apparently very dissimilar to all other members (see below).

Yi et al. (2004) suggest that the basic chromosome number in Apiales is x = 6 (see also Raven 1975), although the order then would have all polyploid/dysploid members, with several independent origins of polyploidy.

Chemistry, Morphology, etc. Kårehed (2003, for which see for further details, also Chandler & Plunkett 2004) provides a good summary of what is known of the main clades in Apiales. For wood anatomy, see Lens et al. (2008a). Endress (2003c) summarizes the rather sparse literature on nucellus development in the clade. For nectary morphology, see Erbar and Leins (2010).

Phylogeny. In earlier studies a grouping [Griselinia [Aralidium + Torricellia]] was rather weakly supported (Backlund
& Bremer 1997; see also Chandler & Plunkett 2002). Plunkett (2001) and Lundberg (2001c) both suggested that Torricelliaceae and Griseliniaceae were successive clades near the base of Apiales (Kårehed 2002a: four genes, all genera sampled, strong Bayesian support), and this topology is followed here (see also Soltis et al. 2011; Tank & Donoghue 2010); the relationship is reversed in Chandler and Plunkett (2004), but with a p.p. of 0.93.

The ex-icacinaceous Pennantia may be sister to all other Apiales, at least in chloroplast phylogenies (e.g. Kårehed 2003; Lens et al. 2008a: strong support), although nuclear markers place it elsewhere in the campanulids (e.g. Chandler & Plunkett 2004). Its position must be confirmed, but the monophyly of the rest of the order is not in doubt.

Pittosporaceae are sister to the rest of
the clade, a clade made up of the old Apiaceae and Araliaceae (e.g. Kårehed 2002c; Andersson et al. 2006: strong support, Myodocarpaceae not included), a position that was strongly supported by Tank and Donoghue (2010). However, some earlier studies found Pittosporaceae to be embedded in the clade, some Bayesian analyses giving strong posterior probabilities for a relationship between Myodocarpaceae and Pittosporaceae (Chandler & Plunkett 2004), or they provided only weak support for a sister group position (Nicolas & Plunkett 2009). Overall, the relationships [Pittosporaceae [Araliaceae [Myodocarpaceae + Apiaceae]]] seem most likely (Tank et al. 2007; Tank & Donoghue 2010; Soltis et al. 2011; Nicolas & Plunkett 2014).

There are some similarities between the [Pittosporaceae [Araliaceae [Myodocarpaceae + Apiaceae]]] clade and Asterales like Campanulaceae and Asteraceae (e.g. Erbar & Leins 2004; Leins & Erbar 2004b), thus 1-benzyltetrahydroisoquinoline alkaloids are found in both (Kubitzki et al. 2011). However, given the rather strongly supported set of relationships at the base of Apiales, the other clades that are in this part of the campanulids, and relationships within Asterales (Campanulaceae and Asteraceae are not immediately related), most of these similarities are parallelisms.

Chemistry, Morphology, etc. Collenchyma is only poorly developed, and a
pericyclic sheath is present; pits in general are bordered. For the morphology of the stigma, see Kårehed (2002b). For information, which should be confirmed, on ovule morphology, see Mauritzon (1936c).

Chemistry, Morphology, etc. Polyacetylenes, mainly aliphatic, including the C17 acetylenes, falcarinone, etc., are found in the [Pittosporaceae [Araliaceae [Myodocarpaceae + Apiaceae]]] clade, while the polyacteylenes of Torricellia angulata, quite recently described, are C11 acids that are unique in having a chiral center (Pan et al. 2006); iridoids are found in the same species (Liang et al. 2009).

For the wood anatomy of this clade, see Noshiro and Baas (1998) and Lens et al. (2008a).

Age. The age of this node is estimated to be (81-)53(-27) m.y. (Lemaire et al. 2011b) or ca 57.2 m.y.a. (Nicolas & Plunkett 2014).

Evolution.Divergence & Distribution. For the fossil history of the East Asian endemic Torricellia, see Meller (2006), Manchester et al. (2009) and Collinson et al. (2012); the genus was widespread in the northern hemisphere by the Eocene and can be dated to a minimum of 55.8 m.y. ago.

Chemistry, Morphology, etc. For fruit and seed of Melanophylla,
see Trifonova (1998), for a revision, see Schatz et al. (1998). Other information
is taken from from Philipson (1977), Lobreau-Callen (1977: pollen), Philipson and Stone (1980), and Takhtajan (2000).

Phylogeny. For the circumscription of this clade, see also Plunkett et al. (2004c); relationships are [Aralidium [Melanophylla + Torricellia]] (see also Soltis et al. 2011).

Previous Relationships.Melanophylla was included in
Hydrangeales and Torricellia and Aralidium were placed in separate monogeneric orders, both in
Cornidae-Cornanae, by Takhtajan (1997); all have been placed in Cornaceae s.l. in the past.

Age. Naumann et al. (2013) suggested an age of a mere 33.9 m.y.a. or so for this node; (115.8-)103.1(-90.2) m.y.a., three times as old, is the estimate in Nicolas and Plunkett (2014), while Magallón et al. (2015), at around 70.5 m.y.a., close to split the difference.

Evolution.Divergence & Distribution. For the various interpretations of the dijunct austral distribution of the family, see Nicolas and Plunkett (2014); in particular, given the relative youth of the crown group, bird-assisted movement from Australia/New Zealand to South America seems most plausible.

For general information, see Philipson (1977) and Dillon and Muñoz-Schick (1993), for seed fatty acids, see Badami and Patil (1981), in part, for leaf
insertion, c.f. Philipson (1967), and for leaf base, c.f. Takhtajan (1997); Takhtajan (2000) provides details
of embryo and testa, for flowers, see Eyde
(1964), and for the ovule, see Warming (1913).

Previous Relationships. Eyde (1964) suggested
relationships of Griselinia to Garrya (Garryales) and Cornaceae (Cornales); the genus was placed in a monogeneric order in
Cornidae-Cornanae by Takhtajan (1997).

Evolution.Divergence & Distribution. Australasia seems the most likely place of origin of the whole clade, and of Pittosporaceae, Araliaceae, Myodocarpaceae and Apiaceae (Nicolas & Plunkett 2014).

Plant-Animal Interactions. Ehrlich and Raven (1964) noted that some butterflies that do not seem to like Araliaceae, including Hydrocotyle, are found on Apiaceae (?including Saniculoideae); thus the Papilio machaon group is not found on Araliaceae because that family lacks the furanocoumarins that the caterpillars like (Berenbaum 1983).

Genes & Genomes. For the complex history of the RPB2 gene (DNA-dependent RNA polymerase) duplications, see Nicolas and Plunkett (2013).

Chemistry, Morphology, etc. There are a number of characters that may delimit clades of various sizes here. These include the presence of sesquiterpene lactones and
benzylisoquinoline alkaloids; presence of crystal sand.

Triterpenoid ethereal oils produce
the distinctive odour characteristic of many of these plants; see Jay (1969)
for suggestions based on plant chemistry that members of this group are related. Kleiman and Spencer (1982) surveyed Apiaceae and Araliaceae for the occurrence of petroselenic acid. Triterpenoid saponins like oleanene are found throughout the group, and also elsewhere (Wang et al. 2012). Lateral roots originate from either side of the xylem poles because a resin canal runs down the stem at the apex of the pole (van Tieghem & Douliot 1888). The vessel:ray pits are bordered
(Baas et al. 2000).

Members of both Araliaceae and Apiaceae in clades that are sister to the rest of these families have simple leaves, as have Myodocarpaceae (Plunkett 2001) and of course Pittosporaceae, so compound leaves may have evolved independently in Apiaceae and Araliaceae. Leaf teeth have a broad glandular
apex with a main and two accessory veins, or one vein proceeds above the
tooth; a survey of tooth morphology might be interesting.

Some Pittosporaceae and a few Araliaceae have basally connate petals (Plunkett 2001). For information on ventral carpel bundles, see Philipson (1970) and Eyde and
Tseng (1971); when the united ventral bundles are opposite the carpels, the two
bundles come from the same carpel, and when they are in the septal radii, the
two bundles are derived from adjacent carpels. For ovule morphology, see Jurica (1922) and van Tieghem (1898). According to the latter, Apiaceae have a thick integument (e.g. ca 7 cells - Gupta 1970), that of Hedera is very thick, although Philipson (1977) described the integument of Araliaceae as
being "thin". There is variation both in the kinds of calcium oxalate crystals that are found in the fruit wall, i.e., single rhomboidal crystals or druses, and also where they occur, e.g. in the endocarp or mesocarp, or all around the fruit or only in commissural area (Liu et al. 2006); I have only just begun to work out the phylogenetic significance of this variation (see also Rompel 1895; Burtt 1991a).

For similarities in wood anatomy between Apiaceae and Araliaceae, see Metcalfe and Chalk (1983); for the sequence of initiation of parts of the flower, see Erbar and Leins (1985: mostly Apiaceae, also Hydrocotyle); for chromosome numbers and evolution, see Yi et al. (2004); for fruit wings and fruit anatomy, see Liu et al. (2006); for nectaries, see Erbar and Leins (2010), and for bark anatomy, see Kotina et al. (2011: crystal types and distributions not integrated into the phylogeny).

Phylogeny. The old woody Araliaceae/herbaceous Apiaceae distinction is no longer tenable, and recent rearrangements have considerable implications for character evolution (c.f. Valcárcel et al. 2014 in part). Myodocarpaceae and Apiaceae-Mackinlayoideae as recognised here both include genera that used to be in Araliaceae, but the precise relationships of the former in particular have been uncertain - see Plunkett and Lowry (2001), Lowry et al. (2001), Plunkett (2001) and Chandler and Plunkett (2004) for more information. Since there are a number of morphological similarities between the Mackinlaya clade and Apiaceae s. str. (Chandler & Plunkett 2002, 2004), Mackinlayoideae are included in Apiaceae here. The old Apiaceae-Hydrocotyloideae, herbaceous and
with simple leaves, are polyphyletic. The large genus
Hydrocotyle is in Araliaceae, a position that makes morphological sense, and although sampling of Hydrocotyle and the related Trachymene must be improved, this is unlikely to affect their position. Arctopus is now in Apiaceae-Saniculoideae; Azorella and a group of genera form a well supported Apiaceae-Azorelloideae, and Centella,
Micropleura, Actinotus, etc., are in Apiaceae-Mackinlayoideae (e.g.
Downie et al. 1998, Downie et al. 2000; Chandler & Plunkett 2003, 2004; Plunkett et al. 2004; Andersson et al. 2006; esp. Nicolas & Plunkett 2009). Nodes along the backbone of this part of the tree have rather moderate support (e.g. Nicolas & Plunkett 2009), yet very few genera remain to be sequenced.

Pittosporaceae are aromatic woody and sometimes thorny plants with simple, estipulate leaves and rather conspicuous flowers that have sepals, petals and stamens equal in number and a gynoecium often with parietal placentation. The fruits are often capsular and enclose pulpy-resinous seeds; the calyx is deciduous.

Evolution.Divergence & Distribution. Many of the apparently plesiomorphic characters of Pittosporaceae, e.g. superior ovary with several ovules, etc., are likely to be derived (see above.

Chemistry, Morphology, etc. Glucosinolates are reported from Bursaria spinosa, but this is probably a mistake (Fahey et al. 2001).

In Pittosporum, at least, the flowers are often functionally unisexual. There is a tendency, especially evident in taxa like Cheiranthera, for the flowers to be obliquely monosymmetric; asymmetry is largely because of the position of the stamens and gynoecium. In Pittosporum floribundum, ovules are epi- and apotropous within a single ovary (Narayana & Sundari 1983). Mauritzon (1939a) drew attention to the fact that the funicular bundle appears to expand and end in the upper part of the funicle, rather than in the chalazal region. This may be connected with how the embryo sac curves; I have not looked for this feature in other Apiales. Seedlings of Pittosporum have up to five cotyledons, and seedling leaves may be pinnatifid.

For embryology, see Narayana and Sundari (1983) and references, for endothelium in particular, see Batygina et
al. (1985), for seed oils (undistinguished), Stuhlfauth et al. (1985), for vegetative anatomy, Wilkinson (1992, 1998), and for testa anatomy, see Takhtajan (2000:
the ovules may be apotropous).

Phylogeny. Cayzer (references in Chandler et al. 2007) has carried out a number of morphological revisions and phylogenetic analyses of parts of Pittosporaceae; changes in taxon limits that she suggests are largely confirmed by a preliminary molecular study (Chandler et al., in Plunkett et al. 2004c). However, generic limits in the Billardiera-Sollya area are still unclear (Chandler et al. 2007). For relationships between species of Pittosporum found in the Pacific, see Gemmill et al. (2002).

Previous relationships. Pittosporaceae were included in Rosales by Cronquist
(1981) and Mabberley (1997). However, evidence has been mounting for over 130 years that they were best associated with Apiaceae/Araliaceae (Hegnauer 1969b for a summary of some literature), van Tieghem (18) even placing Pittosporaceae, Aopiaceae and Araliaceae in his Ombellinées. Takhtajan (1997) included Pittosporaceae as a separate order in his Aralianae (but along with Byblidales).

Age. The age of this node is estimated as (48-)38, 35(-25) m.y. (Bell et al. 2010), while Wikström et al. (2001) thought that this node was (49-)45, 41(-37) m.y. old, Xue et al. (2012) (43.6-)41.1(-40.4) m.y., and Magallón et al. (2015) ca 60.2 m.y.; a much older estimate of (106.6-)94.6(-82.6) m.y. was given by Nicolas and Plunkett (2014).

Evolution.Pollination Biology & Seed Dispersal. For the evolution of andromonoecy in this clade, very rare elsewhere in flowering plants and perhaps connected with dichogamy, successively flowering umbels, and umbels as functional units in pollination, see Schlessman (2011). Interfloral dichogany is common in the clase, and although perfect flowers in Araliaceae are protandrous, as might be expected, about 40% of the records for Apiaceae are for protogyny (Bertin & Newman 1993).

Chemistry, Morphology, etc. Mention of the fruits being dorsi-ventrally or laterally compressed refers to the shape of the contents of individual seeds in transverse section of the mericarp. Individual mericarps can appear to be dorsi-ventrally flattened if they have lateral wings, while two non-flattened mericarps that have not separated can appear to be laterally flattened.

For wood and stem anatomy, see Rodriguez C. (1957) and Oskolski (2001), for young stem and petiole anatomy, see Mittal (1961), for the anatomy of the fruit wing, see A. R. Magee et al. (2010a), and for the rpl16 deletion, see Downie et al. (2000a).

Age. The age of crown Araliaceae is estimated to be (100-)80(-70) m.y. (Mitchell et al. 2012); Beaulieu et al. (2013a: 95% HPD) suggested an age of (48-)44(-41) m.y. and an age of (83.2-)65(-48.7) m.y. has also been mentioned (Nicolas & Plunkett 2014).

Age. For the clade [Gastonia + Hedera] an age of around 85 m.y. or more can be inferred from Valcárcel et al. (2014), while ages of (100-)84(-70) m.y. are suggested for the [Schefflera longipedicellata + The Rest] clade (Mitchell et al. 2012), while the clade [Harmsiopanax + The Rest] was estimated to be around 57.7 m.y.o. (Nicolas & Plunkett 2014) - but c.f. the ages above for the family.

Synonymy: Hederaceae Giseke, Botryodendraceae J. Agardh

Araliaceae may be recognised quite easily. They are often rather stout-stemmed and
little-branched shrubs or trees, rarely herbs, with large and prominent scars
from the fallen leaves; the plant may smell strongly. The leaves are often variously compound and with broad bases,
although these latter rarely more or less encircle the stem; stipules are
common and are morphologically diverse, although they are generally borne on
the leaf base. Petioles are often long, and/or they vary considerably in length on the one shoot; the leaflets are commonly articulated. The ultimate units of
the inflorescence are umbels or heads, the flowers are often rather small,
there are often three or more carpels and the petals are often valvate; the fruit is
drupaceous, sometimes flattened laterally, and the seeds may be ruminate.

Evolution.Divergence & Distribution. Major groupings within Aralioideae in particular show some geographical signal (see below), even if they do not map on to previous classifications. See Mitchell et al. (2012) for some dates in the Greater Raukaua clade, Valcárcel et al. (2014) for dates in the Asian Palmate clade, which depend on the marker used, and Nicolas and Plunkett (2014) for some dates from alomg the spine of the subfamily.

Genes & Genomes. For shifts in the rate of molecular evolution within Araliaceae that are correlated with changes in habit, see Smith and Donoghue (2008).

Chemistry, Morphology, etc.Hedera may have an interrupted
fibrous pericyclic sheath; creeping forms have two-ranked leaves. As to stipules: Fatsia
(no stipules) x Hedera (no stipules) = XFatshedera (stipules),
but some species of Hedera do have a hollowed leaf base with a
flanged margin that encloses the axillary bud.

Araliaceae show considerable floral variation, and this is reflected in their floral vasculature. The calyx may be entirely absent (e.g. Hydrocotyle: Tseng 1967), not even reduced vascular traces suggesting that it was ever there, the petals may have three traces and may be slightly to completely connate (Osmoxylon has basally connate petals), and the stamens sometimes have two traces (Nuraliev et al. 2010, 2011). Both Aralioideae and Hydrocotyle
show early corolla tube initiation (Leins & Erbar 1997; Erbar & Leins 2004). Tetraplasandra gymnocarpa and T. kavaiensis
have secondarily more or less completely superior ovaries (Costello & Motley 2000, 2001, 2004 - see photograph on the cover of American J. Bot. 91(6). 2004). Hydrocotyle, almost alone in the family, is highly polyploid (Yi et al. 2004).

Variation in meristicity is particularly striking in Aralioideae, and flowers are up to 12-merous or more there (Viguier 1906). Tupidanthus calyptratus (= Asian Schefflera) has up to 172 stamens and 132 carpels; the carpels are initiated in a single elongated and sometimes contorted whorl looking rather like a brain cactus (fasciation: Oskolski et al. 2005; Nuraliev et al. 2009, 2014; see also Eyde & Tseng 1971). Some species have about as many stamens as carpels, separate members of the calyx cannot be distinguished (the calyx forms a low, entire rim; Nuraliev et al. 2014), the petals may be connate and form a calyptra, and although the symplicate zone of the gynoecium develops first, the carpels are largely synascidiate; there is a large, flat remnant of the floral axis within the carpel whorl (Sokoloff et al. 2007b). Plerandra has up to 25 or more carpels and to 500 stamens, while other species there are up to five series of stamens initiated centripetally, the vasculature of members of each whorl being connected radially (very unusual for euasterids); carpel number is 14 or fewer (Philipson 1970; Oskolski et al. 2010c). For further discussion, see the [euasterid] clade.

For general information, see Philipson (1970), for general anatomy, see Viguier (1906), for that of the leaf, see de Villiers et al. (2010), for bark, see Kotina & Oskolski (2010), for wood, see Oskolski (1996) and Kotina et al. 2013 and references, for gynoecial development in Seemannaralia, see Oskolski et al. (2010b), for embryology, see Ducamp (1902), Gopinath (1944) and Mohana Rao (1973b), and for fruit anatomy, see Konstaninova and Suchorukow (2010). For Hydrocotyle: see Leins and Erbar (2010), flowers; Shu and She (2001), pollen of Chinese spp; Håkansson (1923), ovules.

Phylogeny. Basal Araliaceae may well be bicarpellate
(see also Wen et al. 2001) and have simple leaves, as in the herbaceous Hydrocotyloideae (ex
Apiaceae), which are sister to the rest of the family (Chandler & Plunkett 2004; Plunkett et al. 2004a; Nicolas & Plunkett 2009). The S.W. Australian genera Neosciadium and probably Homalosciadium also belong to Hydrocotyloideae (Andersson et al. 2006). Hydrocotyle has laterally flattened fruits with a sclerified (= woody) endocarp and stipules that are either cauline or are borne on the leaf base - and it also has trilacunar nodes (Sinnott & Bailey 1914). Trachymene, perhaps to include Uldinia, is also in Hydrocotyloideae; morphologically it is rather similar to them, and although there is a carpophore in the fruit, it is undivided. For a phyogeny of Trachymene, see Henwood et al. (2010).

Astrotricha and Osmoxylon may be part of a polytomy at the node immediately above Hydrocotyloideae (see also Lee et al. 2008), while the position of Harmisopanax, which has fruits that are schizocarpic like those of Hydrocotyloideae, is also uncertain (Nicolas & Plunkett 2009).

For more details on the phylogeny of Aralioideae in particular, see Henwood and Hart (2001) and especially Wen et al. (2001), Plunkett et al. (2004a, c), and Lowry et al. (2004). Schefflera, with perhaps 1,600 species under 2/5 of which have been described (Frodin et al. 2010), is highly polyphyletic, and five major clades have become apparent in it - of which Schefflera s. str. is perhaps the smallest. These are circumscribed geographically and some also have morphological support. African plus Madagascan taxa form a clade, as do the some 250-300 neotropical species, the Asian species (the last two clades are close on the tree), and two groups of species restricted to the Pacific, the small Schefflera s. str. and the much larger Melanesian clades (Plunkett et al. 2005 [general], 2009 [Melanesia], 2010 [Neotropics]; Gostel et al. 2009 [Africa-Madagascar]; Fiaschi & Plunkett 2011 [Neotropics]; Plunkett & Lowry 2012 [Melanesia]: Frodin et al. 2010 for a summary). In part these clades map on to earlier infrageneric groupings, although with the inclusion of segregate genera.

The Schefflera problem aside, there are four major clades in Aralioideae, the largely South East Asian Palmate and Aralia-Panax groups, and the Pacific and Indian Ocean basin Greater Raukaua (Mitchell et al. 2012) and the Polyscias-Psuedopanax groups (Wen et al. 2001; Mitchell & Wen 2004; Plunkett et al. 2004a; see Valcárcel et al. 2014 for a summary). Within the Palmate group, which includes Hedera, relationships are rather poorly resolved, and details depend on the markers used - polyploidy and ancient hybridization may be involved - and the position of Osmoxylon seems particularly uncertain (Yi et al. 2004; Mitchell & Wen 2004; Valcárcel et al. 2014). Lee et al. (2008) focused on relationships of Malesian Araliaceae; Osmoxylon was isolated. Most (?all) Dendropanax are likely to be monophyletic, with two large clades restricted to the Old and New Worlds (Li & Wen 2013).

Classification. For a now dated checklist of the family, see Frodin and Govaerts (2003), although this is already very dated. Generic limits in Aralioideae need much attention; see Plunkett et al. (2004a) for genera, but more changes will be needed. The Pacific clade of Schefflera is to be called Plerandra; Polyscias has been substantially enlarged (Lowry & Plunkett 2010).

Chemistry, Morphology, etc. In wood anatomy Myodocarpus is perhaps more like Cornaceae than any other members of the Apiaceae-Araliaceae complex, but in other features it is more like Apiaceae (Rodrigues C. 1957). Myodocarpus has a number of
distinctive features of the flower and in particular its schizocarp. The mericarps are beautiful little laterally-flattened samaras that at first sight are similar to those of Serjania.

Apiaceae are usually herbs
with compound leaves that have broad, sheathing bases. Their inflorescences are
usually umbels of umbels, rarely heads; and their polypetalous flowers usually have a
minute calyx, clawed petals with incurved apices, and two carpels; the fruits
are dry and separate into two 1-seeded portions.

Saniculeae have usually undivided leaves with hairs or spiny tips to the teeth, sessile and spiny or scaly fruits, an often apparently more
or less simple umbel or head, and there is no free carpophore.

Apioideae usually have more or less finely-divided leaves, compound umbels, and the usually pedicellate fruit has a free,
bifid carpophore.

Evolution.Divergence & Distribution. As relationships get sorted out, evolutionary and biogeographic studies become possible. Nicolas and Plunkett (2014, q.v. for details) suggested an Australian origin for Apiaceae, with at least some scenarios suggesting a South American origin for Hydrocotyloideae and an African origin for Apioideae and Saniculoideae. Indeed, a glance at the maps of the clades from Hermas to core apioids shows a concentration of taxa in southern Africa in particular, the three basal clades of Apioideae and the two basal clades in Saniculoideae being small and made up very largely of southern African genera (see also Banasiak et al. 2013; Nicolas & Plunkett 2014; this also has considerable implications for character optimisations (Calviño et al. 2006; Magee et al. 2010a for details).

A number of taxa in the small clades in "basal" Apiaceae are woody and have undivided leaves (see the characterisations above). Although these woody Apiaceae are quite common, the ancestral habit for Apioideae, at least, may be herbaceous (Calviño et al. 2006). Woodiness in Bupleurum is derived and has evolved several times (H.-C. Wang et al. 2013); Myrrhidendron, from Central America and Colombia, and a few other euapioids, are also woody. The place of origin of Bupleurum may be in archipelagic early Caenozoic Europe, with much subsequent diversification in eastern Asia, although anywhere from Europe and North Africa to Eastern Asia may have been involved (Banasiak et al. 2013, q.v. for much more detail).

Spalik et al. (2010) looked at wide disjunctions in Apioideae, providing dates for various nodes, and later (Spalik et al. 2014) focussed on biogeographic patterns in the largely aquatic Oenantheae where there seems to have been much dispersal from Eurasia to North America. Spalik et al. (2014) found that the Hawaiian "Peucedanum sandwicense" (a split from stem Oenanthe) may have arrived on Hawaii around 17.2 m.y.a., so being another example of a clade older than the current islands. Lilaeopsis brasiliensis and L. mauritiana are very close, even though they are separated by the Atlantic Ocean and the African continent (Spalik et al. 2010).

Carpophores, chromosome numbers other than 8, and inflorescences that are not condensed (all in African taxa) could be plesiomorphic in Saniculoideae; simple umbels are apomorphic for Saniculeae, etc. (Magee et al. 2010a, q.v. for numerous character optimisations; see also Calviño et al. 2008a; Kadereit et al. 2008).

Plant-Animal Interactions. Caterpillars of Papilionidae-Papilionini butterflies are notably common (ca 13% of all records) on Apiaceae, perhaps shifting here from Aristolochiaceae (Fordyce 2010; see also Berenbaum & Feeny 2008; Simonsen et al. 2011; Condamine et al. 2011). Interestingly, they are not found on either Pittosporaceae or Araliaceae; thus they will not eat Hydrocotyle (here Araliaceae), many of the larvae of Papilio ajax tested absolutely refusing to eat it (see also Dethier 1941; Ehrlich & Raven 1967). Linear fumarocoumarins are often phototoxic but are tolerated by caterpillars that will not eat plants with the non-photoxic linear coumarins (Berenbaum & Feeney 1981). Microlepidopteran larvae of the Elachistidae-Depressariinae are common on Apiaceae, although they may have initially been associated with rosids and have also colonized some Asteraceae (Fetz 1994: host plants; Berenbaum & Zangerl 1998: chemistry of the [co]evolution of resistance; Berenbaum & Passoa 1999: phylogeny). There has been a diversification of agromyzid dipteran leaf miners in north temperate Apiaceae; they were previously on Ranunculaceae, also a group with noxious secondary metabolites (Winkler et al. 2009). For general insect-umbellifer relationships, see Berenbaum (1990) and Sperling and Feeny (1995).

Pollination Biology & Seed Dispersal. All the flowers in an umbel open more or less simultaneously. In a number of Saniculoideae and a few Hydrocotyloideae in particular the inflorescence bracts function as petals, the rest of the inflorescence being much reduced and the whole looking more or less like a simple flower (Froebe & Ulbrich 1978). In the umbels of a number of Apioideae, the marginal flowers are larger than the others, and their abaxial petals may be larger than the adaxial, so increasing the resemblance of the inflorescence to a polysymmetric flower. The dark flower in the centre of the umbel of taxa like Daucus carota may attract flies that pollinate the flowers (Westmoreland & Muntan 1996).

Although the flowers appear unspecialized, being potentially pollinated by a variety of pollinators, like other such systems, e.g. Asteraceae, oligolectic pollinators may play a major role in pollination - in the case examined, the bee Andrena ziziae as a pollinator of Thaspium and in particular Zizia in the southeast U.S.A. (Lindsey 1984; Lindsey & Bell 1985). However, as with other such systems, other bees (and a syrphid fly) were also effective pollinators, and pollinators varied with geography, too.

Vegetative Variation. Variation in leaf morphology, even within Apioideae, is considerable. Thus the small circum-Pacific genus Oreomyrrhis (= Anthriscus) includes species with ordinary-looking highly dissected leaves, leaves with a series of almost tooth-like leaflets on either side of the rachis, linear leaves, sometimes lobed at the apex, although the lobes are not articulated, and small, undivided leaves. In the last case the plant is tussock-forming and almost moss-like, and I have seen specimens identified as Centrolepidaceae (= Restionaceae), a monocot! Species with all these leaf morphologies are found on the mountains of New Guinea. Although Oreomyrrhis is probably monophyletic, it is well embedded in Chaerophyllum, a genus hitherto thought to be fairly well understood (Chung et al. 2005; Chung 2007). Lilaeopsis occidentalis and Oxyopolis greenmanii have linear, terete leaves that are marked by articulations at intervals. These leaves are comparable with the rhachis of compound leaves, hydathodes borne at the articulations representing much reduced and modified pinnae (Kaplan 1970b, c.f. esp. Figs 3E and 6A). Such leaves appear to have evolved several times (Feist & Downie 2008; Feist et al. 2012). Bupleurum has undivided leaves, those of B. rotundifolium being almost orbicular, entire, and perfoliate, hence its common name, thorow wax.

Seeds with relatively longer embryos may have evolved in open habitats and in plants with an annual life cycle (Vandelook et al. 2012b). There is considerable variation in seedling morphology. A number of taxa have cryptogeal germination during which the plumule ends up, planted, as it were, under ground; associated with this, monocotyly is also quite common (see Haccius 1952b; Cerceau-Larrival 1962; Haines & Lye 1979).

Genes & Genomes. For the occurence of the coxl pseudogene (a mitochondrial gene) in the plastid in Daucus and relatives, see Straub et al. (2013).

Chemistry, Morphology, etc. Gums and resins are scattered in the family. In Apiaceae the flavone apigenin is synthesized by an enzyme belonging to the oxoglutarate dependen dioxygenase family, not by a member of the cytochrome P450 family, as is usual in angiosperms (Pichersky & Lewinsohn 2011). For the distinctive rosmarinic acid glucoside found in Saniculoideae-Saniculeae, see Olivier et al. (2008). Oskolski et al. (2010a) describe wood and bark anatomy of Steganotaenieae in particular and Saniculoideae in general. Peripheral collenchyma in
the stem is often especially well developed, and petiole anatomy is very
variable and complex (e.g. Metcalfe 1950). Taxa such as Foeniculum have stipules of sorts.

The usually compact inflorescence units of Saniculoideae may be best interpreted as a group of reduced umbellules (Froebe 1964, 1971), although they are called simple umbels above. Centella (Mackinlayoideae) has a few branches from the petal bundle (Gustafsson 1995); petal vasculature may repay attention. Spichiger et al. (2002) show the two carpels as being collateral. According to Eyde and Tseng (1971), whether the ventral carpel bundles are fused bundles of adjacent placentae or are from the same placenta varies within Apiaceae without any particular systematic significance. Van Tieghem (1898) noted that in Apiaceae the ascending ovule aborts and the pendulous ovule persists, however, other reports suggest that both ovules are pendulous (Philipson 1970).

Fruit anatomy is currently being studied by M. Liu and co-workers, and there is a considerable amount of variation which shows at least some correlation with clades. For vittae, druses, etc. in the fruits, see Liu et al. (2007, 2012b), for fruit anatomy of Azorelloideae, see Liu et al. (2009), and for carpophores, see Liu et al. (2012a) - although Phlyctidocarpa (Saniculoideae) by one definition in this last paper would seem to lack a carpophore (group A), yet it is scored as having a bifid carpophore.

For chemistry, see Hegnauer (1971), Berenbaum (2001) and Olivier and van Wyk (2013: Saniculeae), for stomata, see Guyot 1971 and references - value slight?), for wood anatomy in Apioideae-Heteromorpheae and Mackinlayoideae, see Oskolski and van Wyk (2008, 2010) and in woody Saniculoideae, see Oskolski et al. (2010a), for inflorescences of Saniculoideae and Hydrocotyloideae in particular, see Froebe (1964, 1979), for those of Eryngium, see Harris (1999), for floral development, see Leins and Erbar (2004b), for pollen of Chinese Apiaceae, see Shu and She (2001), for the distribution of the hypostase, see Gupta (1970), for embryo sac morphology and fruit anatomy, esp. of genera in the old Hydrocotyloideae, see Tseng (1967), for ovules, etc., see Håkansson (1923), and for chromosome number and morphology, see Pimenov et al. (2003). There is much useful information in Chandler and Plunkett (2003, 2004) and also in all four numbers of Plant Divers. Evol. 128. 2010, inc. Stepanova & Oskolski (2010: Bupleurum); see also Burtt (1991a: southern African genera) and Van Wyk et al. (2013: African taxa) for information about Apiaceae in a biogeographically/phylogenetically critical area.

Phylogeny. The old Apiaceae-Hydrocotyloideae are hopelessly polyphyletic, and their members now occur in several quite separate clades of which most are in Apiaceae (Nicolas & Plunkett 2009). Some genera of Mackinlayoideae used to be in Araliaceae-Mackinlayeae, others in Apiaceae-Hydrocotyloideae. Genera to be included here include Apiopetalum, Mackinlaya, Micropleura and Xanthosia. Melikian and Konstantinova (2006) thought that the gynoecial structure of Actinotus was so different from that of other Apiaceae that the genus deserved to be placed in its own family; it belongs here, although its exact position is unclear (Nicolas & Plunkett 2009).

The Australian Platysace - perhaps to include Homalosciadium - is not a member of Mackinlayoideae, where it had been placed (e.g. Chandler & Plunkett 2004). It is sister to Apioideae in some analyses (Henwood & Hart 2001; Andersson et al. 2006), but a position rather deep in the tree sister to [Azorelloideae [Saniculoideae + Apioideae]] is strongly supported (Nicolas & Plunkett 2009, 2014), although Soltis et al. (2011: but sampling) found that it was moderately supported as sister to Mackinlaya. The positions of Klotzschia ("distinctive fruits") and Hermas, both of which used to be in Hydrocotyloideae, are unclear (Andersson et al. 2006; Calviño et al. 2006, 2008). Klotzschia, herbs to subshrubs with peltate leaves from Brazil, may go with Azorelloideae, Apioideae, or join the back-bone immediately above Azorelloideae (Nicolas & Plunkett 2014). Hermas, a South African endemic, has some similarities with Saniculoideae, but it is unlikely to be placed within any currently recognized subfamily (Nicolas & Plunkett 2009); there is some support for a position as sister to [Saniculoideae + Apioideae] (e.g. Nicolas & Plunkett 2014), and I have tentatively placed it there.

For the phylogeny of Azorelloideae, which includes about half the genera that used to be in Hydrocotyloideae, see Downie et al.
(1998, 2000a, 2001), Mitchell et al. (1999), Plunkett & Lowry (2001), Henwood and Hart (2001: the Bowlesia clade), Chandler and Plunkett (2004), Andersson et al. (2006) and Nicolas and Plunkett (2009: q.v. for details). Stilbocarpa used to be in Araliaceae, but it is probably sister to the Andean Huanaca. Klotzschia may also belong here, but its position is unstable (see above); this genus aside, the South American Diposis is sister to other Azorelloideae (Nicolas & Plunkett 2009). Azorella is para/polyphyletic with respect to Laretia, Mulinum, and three other genera, in particular, the type of Azorella is rather distant from most of the rest of the genus (Nicolas & Plunkett 2012). Bowlesia lacks petroselenic acid, but it was apparently the only Azorelloideae studied (Kleiman & Spencer 1982).

The monophyly
of Saniculoideae (except Lagoecia, now in Apioideae) is upheld
in all molecular analyses, and some details of relationships within it were suggested by Valiejo-Roman et al. (2002). In some analyses the African ex-hydrocotyloid Arctopus (see Magin 1980 for floral development)
is sister to other Saniculoideae, and the woody Steganotaenia and Polemanniopsis, and perhaps Lichtensteinia, are part of the same clade (Downie et al. 2001; van Wyck 2001; M. Liu et al. 2003; Plunkett et al. 2004c; Magee et al. 2010a); at least some of the latter genera have a slightly lignified endocarp, apparently alone in both Saniculoideae and Apioideae (M. Liu et al. 2004). Calviño et al. (2006, esp. 2007: good general discussion of variation), however, expressed reservations about this expanded - and characterless - Eryngioideae, but Calviño and Downie (2007, c.f. Magee et al. 2010a) found that the clade could be circumscribed satisfactorily so long as Lichtensteinia moved to Apioideae; they recognised two tribes, both well supported and with unique indels. A clade including Lichtensteinia and Choritaenia were weakly supported as being sister to other Saniculoideae (Nicolas & Plunkett 2009); although this set of relationships has not been upheld, the monotypic African Phlyctidocarpa does seem to be a member of Saniculoideae, although its exact position there is unclear (Magee et al. 2010a, see also Nicolas & Plunkett 2014). The large genus Eryngium has separate New and Old World clades (Calviño et al. 2008b, 2010); a recent morphological analysis suggested that four unrelated species (in the system later used in the same paper) made up a series of basal pectinations, although there was no strong support for this topology (Wörz 2011).

Relationships at the base of Apioideae are very pectinate and are something like [Lichtensteinia [Annesorhiza clade [Heteromorpheae [Bupleurum + The Rest]]]] (Downie & Katz-Downie 1999; Plunkett et al. 2004; Calviño et al. 2006; Magee et al. 2008, also a revision of Ezosciadium; Magee et al. 2010a; Downie et al. 2010; Nicolas & Plunkett 2014). Apart from the position of Lichtensteinia, in some analyses sister to [Saniculoideae + Apioideae], relationships in Nicolas and Plunkett (2009) are similar; above Bupleurum on the tree was the Mexican Neogoezia. In some analyses the Annesorhiza clade appeared to be allied with Heteromorpheae; Bupleurum is strongly supported as sister to the remainder of the subfamily (Calviño et al. 2005, 2006). For relationships within Bupleurum, in which the sole South African species is derived and a Mediterranean clade of sometimes shrubby, pinnately-veined taxa is sister to the rest of the genus, see Neves and Watson (2004) and H.-C. Wang et al. (2014).

Classification. The subfamilial classification of Plunkett et al. (2004c) and Downie et al. (2010) are largely followed here, but some changes may be needed given the positions taken by some ex Hydrocotyloideae in the trees recovered by Nicolas and Plunkett (2009, 2014). Classical generic and tribal limits in Apioideae are
a notable disaster area. The
traditional tribal and generic classification was based primarily on gross fruit morphology; convergence of fruit
architecture as adaptations to modes of fruit dispersal has resulted in many non-monophyletic
groupings. Downie et al. (2010) placed the 41 major clades they obtained in Apioideae in 30 groups, recognising tribes and subtribes where they exist, and listing included genera; some groups of genera were unnamed. For tribes in Saniculoideae and in the basal pectinations in Apioideae, see Magee et al. (2010a); five of these tribes were newly described (and all are small) and Marlothielleae and Choritaenieae, both monospecific, are combined with Lichtensteinieae above; even an expanded Lichtensteinieae can still muster a mere 9 species.

For genera, old style, see Pimenov and Leonov (1993). Just about all the papers dealing with phylogenetic relationships in Apioideae (see above) have implications for generic limits. For instance, 13/18 genera for which two or more sequences were included in a study by Downie et al. (2000b) were found not to be monophyletic, while Weitzel et al. (2014) included five genera in Thapsia, "the deadly carrot", a genus of under 50 species. Downie et al. (2010) listed 18 genera that were wildly polyphyletic, i.e., had members in two or more of their 41 major clades; Pimpinella was no. 1 in the list, species being known from seven of these clades. The problem is further compounded by the disproportionately large number of mono- or di-typic
genera (see also Spalik et al. 2001; Valiejo-Roman et al. 2006; Spalik & Downie 2007). It is not that fruit characters never correlate with redrawn generic boundaries (c.f. Feist et al. 2012: rhachis-leaved North American taxa; Liu et al. 2012), but they often mislead when used by themselves and/or without detailed examination of fruit anatomy.